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Dunn's Salamander Project

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Dunn's Salamander Project Draft for Review Only – Not for Distribution 1 5 June 2008 2 Marc P. Hayes 3 Habitat Program, Science Division 4 Washington Department of Fish and Wildlife 5 600 Capitol Way N., Mailstop 43143 6 Olympia, WA 98501-1091 7 360/902-2567; Fax: 360/902-2946 8 [email protected] 9 RH: Hayes et al.• Wood and Terrestrial Salamanders 10 Terrestrial Salamander Wood Utilization in Managed Landscapes: Implications for Forestry 11 Practices 12 MARC P. HAYES1, Washington Department of Fish and Wildlife, Habitat Program, 600 Capitol 13 Way N., Olympia, WA 98501, USA 14 TIMOTHY QUINN, Washington Department of Fish and Wildlife, Habitat Program, 600 Capitol 15 Way N., Olympia, WA 98501, USA 16 TIFFANY L. HICKS, Washington Department of Fish and Wildlife, Habitat Program, 600 17 Capitol Way N., Olympia, WA 98501, USA 18 AIMEE P. MCINTYRE, Washington Department of Fish and Wildlife, Habitat Program, 600 19 Capitol Way N., Olympia, WA 98501, USA 20 MARTIN G. RAPHAEL, Pacific Northwest Research Station, Olympia Forestry Sciences 21 Laboratory, 3625 93rd Avenue S.W., Olympia, WA 98512, USA 22 JAMES G. MACCRACKEN, Longview Timberlands, P.O. Box 667, Longview, WA 98632, 23 USA 24 M. ANTHONY MELCHIORS, Weyerhaeuser Company, 32901 Weyerhaeuser Way S., Federal 25 Way, WA 98063, USA 1 Email: [email protected] Draft for Review Only – Not for Distribution 2 | Hayes et al. 26 ANGELA B. STRINGER, The Campbell Group LLC, One S.W. Columbia Street, Suite 1700, 27 Portland, OR 97258, USA 28 ABSTRACT We studied the 4 species of terrestrial salamander in managed forests of 29 southwestern Washington to better understand their use of dead wood as habitat. During April- 30 June, we intensively sampled 10 2-m wide belts oriented perpendicular to stream axes at 14 31 streams in 2001 and 5 streams in 2003. We partitioned belts into macrohabitats (banks and 32 uplands), and characterized habitat within each belt and microhabitats around each salamander. 33 All 4 species were associated with wood, but at different levels, a pattern that seemed related to 34 both spatial (macrohabitat) and temporal (year) patterns in temperature and moisture. Dunn’s and 35 Van Dyke’s salamanders (Plethodon dunni and P. vandykei), the 2 more hydrophilic species, 36 occurred most frequently in the wetter macrohabitat (banks) regardless of year, but also used 37 locally wetter and (in the case of Dunn's salamander) cooler microhabitats during the drier 38 (2001) but not the wetter (2003) year. Though Dunn’s salamander used wood as cover less 39 frequently than Van Dyke’s salamander, its use of wood increased 2.5 times in the drier versus 40 wetter year. Van Dyke’s salamander stood out from the other species in its greater use of large 41 wood, though data were only available for the dry year. In contrast, the 2 less hydrophilic 42 species, ensatina (Ensatina eschscholtzii) and western red-backed salamander (P. vehiculum) 43 were widespread across macrohabitats; and though they exhibited no consistent change in their 44 use of wood under drier conditions, they both selected locally cooler and (in the case of the 45 western red-backed salamander) wetter microhabitats in the drier year. Wood use by terrestrial 46 amphibians is clearly species-specific and an association between the more hydrophilic species 47 and wood, that appears most apparent in dry years, merits further investigation. If Van Dyke’s Draft for Review Only – Not for Distribution 3 | Hayes et al. 48 salamander is dependent on large wood, it may be susceptible to practices that reduce the quality 49 (size) and quantity of that wood, particularly on streambanks. 50 KEY WORDS coarse woody debris, Ensatina eschscholtzii, habitat use, managed landscapes, 51 Plethodon dunni, Plethodon vandykei, Plethodon vehiculum, terrestrial salamanders, timber 52 harvest, Washington. 53 The Journal of Wildlife Management: 00(0): 000-000, 200X 54 Wood debris is believed to be an important resource for many forest-dwelling organisms 55 (Bunnell et al. 1999, Ódor et al. 2001, Bull 2002, Bunnell et al. 2002, Grove 2002, Mathieu et al. 56 2005), including amphibians (Aubry et al. 1988, Aubry and Hall 1991, Corn and Bury 1991, 57 Gilbert and Allwine 1991, Walnick 1997, Butts and McComb 2000). This belief has been based 58 on the frequency with which amphibians were associated with logs (e.g., Corn and Bury 1991), 59 correlations of amphibian abundance with the volume of coarse wood debris (e.g., Butts and 60 McComb 2000), or more limited data on nest site occurrence in downed wood (Hanlin et al. 61 1978, Norman and Norman 1980, Norman 1986, Jones 1989, Blessing et al. 1999, Nauman et al. 62 1999, Olson et al. 2006). Yet, recent work in managed forest suggested that abundance of 63 amphibians was unrelated to the amount of coarse wood debris (Aubry 2000) and adds to the 64 increasing number of studies that demonstrate conflicting results about the importance of wood 65 to terrestrial salamanders (compare Dupuis et al. 1995, Dupuis 1997, Dupuis and Bunnell 1999, 66 and Grialou et al. 2000 to Corn and Bury 1991 and Aubry 2000). 67 We envision 3 major biological reasons why a particular study would fail to show that 68 wood is important to terrestrial amphibians: 1) wood is not important habitat to amphibians and 69 hence, has little effect on suitability of their habitat; 2) wood becomes increasingly important as 70 other types of physical habitat become more limited; and 3) wood becomes important only under Draft for Review Only – Not for Distribution 4 | Hayes et al. 71 certain climatic conditions. The first reason, while plausible, fails to explain why studies of the 72 same species have reached completely opposite conclusions about the wood-amphibian 73 relationship. The second reason has been the subject of speculation. For example, some have 74 suggested that wood and alternative substrates may substitute for one another (Hagar et al. 1995, 75 Bunnell et al. 1997), but the relative value of different substrates to amphibians, at least in the 76 Pacific Northwest, has garnered little attention. The third reason is based in part on distinctive 77 properties of wood and the life history of terrestrial amphibians in the Pacific Northwest. Wood 78 has a substantial capacity to absorb and retain water (Stamm 1935, Stamm and Loughborough 79 1942, Jayme 1958), and thus may create microhabitats that differ from many other substrates, 80 particularly non-porous rocks. Terrestrial amphibians in the Pacific Northwest are exclusively 81 lungless (plethodontid) salamanders (Jones et al. 2005) and rely almost exclusively on moist 82 skin-based gas exchange (Feder and Burggren 1992). Wood may provide hydric or thermal 83 advantages over alternative substrates (Heatwole 1962, Bunnell et al. 2002). In the Pacific 84 Northwest, where salamanders typically restrict their activities to habitats and seasons where risk 85 of desiccation is minimized (Aubry et al. 1988, Grialou et al. 2000, Aubry 2000), availability of 86 wood may be especially important during the summer dry periods. 87 Our overarching purpose was to provide basic data to clarify the relationship between 88 terrestrial amphibians and wood in managed forest landscapes. Our study was structured around 89 Dunn’s salamander (Plethodon dunni) but we included data on Van Dyke’s salamander (P. 90 vandykei), ensatina (Ensatina eschscholtzii) and western red-backed salamander (P. vehiculum). 91 To address this need, we first describe the habitat relationships among the terrestrial salamanders 92 that occur in southwestern Washington State, focusing on the distribution of animals relative to 93 streams, and then on more fine-scale moisture, temperature and wood use patterns. We identify Draft for Review Only – Not for Distribution 5 | Hayes et al. 94 how terrestrial salamander use of wood varied between habitats and years with different moisture 95 and temperature conditions, and suggest how individual salamander species might be influenced 96 by losses of wood associated with forest management practices. 97 98 STUDY AREA 99 The study area was located in the Willapa Hills of southwestern Washington (Figure 1), 100 which comprise the northern segment of the Coast Ranges physiographic province (Franklin and 101 Dyrness 1988). This region had a complex topography of mostly low hills (maximum elevation 102 948 m); valleys with significant alluvial area occurred in only a few larger streams (e.g., 103 Chehalis, Naselle, and Willapa Rivers; Figure 1). Geology, a complex mix of Tertiary 104 formations, included mostly marine sedimentary (mudstone, siltstones, sandstones) and intrusive 105 basalt formations (Franklin and Dyrness 1988). Over 95 percent of the region was managed for 106 timber. We chose this area because it is the only part of Washington State in which Dunn’s 107 salamander was known to occur (Dvornich et al. 1997). 108 109 METHODS 110 Site Selection 111 We selected sites along 19 streams (Figure 1) that were: i) distributed across the study 112 area; ii) located on first-to-fourth-order streams (Strahler 1952); and that iii) had at least a 120-m 113 reach surrounded by forest stands greater than 15 years in age. These criteria ensured that we 114 sampled a range of stream sizes but none so large that it prevented us from easily crossing the 115 stream to sample both sides. Also, by ensuring that riparian forest was greater than 15 years of 116 age, we were able to avoid stream banks that were inaccessible due to heavy accumulation of Draft for Review Only – Not for Distribution 6 | Hayes et al. 117 harvest debris (Jackson and Sturm 2002; personal observation). Eighteen streams were from 118 forests managed for timber; one was located on unmanaged land. 119 120 Sampling 121 Sampling overview.— We used a streambank survey approach modified from Raphael et 122 al.
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